A hypothesis for the evolution of androdioecy: the joint influence of reproductive assurance and local mate competition in a metapopulation

Akimoto, J., Fukuhara, T. and Kikuzawa, K. (1999) Sex ratios and genetic variation in a functionally androdioecious species, Schizopepon bryoniaefolius (Cucurbitaceae). Am. J. Botany 86, 880-886. Aronne, G. and Wilcock, C.C. (1994) Reproductive characteristics and breeding system of shrubs in the Mediterranean region. Funct. Ecol. 8, 69-76. Ashman, T. (1994) Reproductive allocation in hermaphroditic and female plants of Sidalcea oregana spp. spicata (Malvaceae) using four currencies. Am. J. Botany 81, 433-438. Baker, H.G. (1955) Self-compatibility and establishment after “long-distance” dispersal. Evolution 9, 347-348. Baker, H.G. (1967) Support for Baker's Law — as a rule. Evolution 21, 853-856. Baker, H.G. and Cox, P.A. (1984) Further thoughts on dioecism and islands. Ann. Missouri Bot. Garden 71, 244-253. Barrett, S.C.H. and Pannell, J.R. (1999) Metapopulation dynamics and mating-system evolution in plants. In P. Hollingsworth, R. Bateman and R. Gornall (eds) Molecular Systematics and Plant Evolution. Chapman and Hall, London, pp. 74-100. Barrett, S.C.H. and Shore, J.S. (1987) Variation and evolution of breeding systems in the Turnera ulmifolia complex (Turneraceae). Evolution 41, 340-354. Belaoussoff, S. and Shore, J.S. (1995) Floral correlates and fitness consequences of mating-system variation in Turnera ulmifolia. Evolution 49, 545-556. Burd, M. and Allen, T.F.H. (1988) Sexual allocation strategy in wind-pollinated plants. Evolution 42, 403-407. Charlesworth, D. (1984) Androdioecy and the evolution of dioecy. Biol. J. Linn. Soc. 22, 333-348. Charlesworth, D. and Charlesworth, B. (1978) A model for the evolution of dioecy and gynodioecy. Am. Nat. 112, 975-997. Charlesworth, D. and Charlesworth, B. (1981) Allocation of resources to male and female functions in hermaphrodites. Biol. J. Linn. Soc. 15, 57-74. Charlesworth, D. and Charlesworth, B. (1987) Inbreeding depression and its evolutionary consequences. Ann. Rev. Ecol. Syst. 18, 273-288. Charlesworth, D., Morgan, M.T. and Charlesworth, B. (1990) Inbreeding depression, genetic load, and the evolution of outcrossing rates in a multilocus system with no linkage. Evolution 44, 1469-1489. Charnov, E.L. (1982) The Theory of Sex Allocation. Princeton University Press, Princeton, NJ. Charnov, E.L., Maynard Smith, J. and Bull, J.J. (1976) Why be an hermaphrodite? Nature 263, 125-126. Couvet, D., Ronce, O. and Gliddon, C. (1998) The maintenance of nucleocytoplasmic polymorphism in a metapopulation: the case of gynodioecy. Am. Nat. 152, 59-70. Cruden, R.W. (1977) Pollen-ovule ratios: a conservative index of breeding systems in flowering plants. Evolution 31, 32-46. Cruzan, M.B., Hamrick, J.L., Arnold, M.L. and Bennett, B.D. (1994) Mating system variation in hybridizing Irises — effects of phenology and floral densities on family outcrossing rates. Heredity 72, 95-105. Darwin, C. (1877) The Different Forms of Flowers on Plants of the Same Species. Appleton, New York. Dommee, B., Geslot, A., Thompson, J.D., Reille, M. and Denelle, N. (1999) Androdioecy in the entomophilous tree Fraxinus ornus (Oleaceae). New Phytol. 143, 419-426. Durand, B. (1963) Le complexe Mercurialis annua L. s.l.: une etude biosystematique. Annales des Sciences Naturelles, Botanique, Paris 12, 579-736. Durand, R. and Durand, B. (1992) Dioecy, monoecy, polyploidy and speciation in annual Mercuries. Bull de la Societé Botanique de France, Lettres Botaniques 139, 377-399. Farris, M.A. and Mitton, J.W. (1984) Population density, outcrossing rate, and heterozygous superiority in Ponderosa pine. Evolution 38, 1151-1154. Fritsch, P. and Rieseberg, L.H. (1992) High outcrossing rates maintain male and hermaphrodite individuals in populations of the flowering plant Datisca glomerata. Nature 359, 633-636. Gouyon, P.H. and Couvet, D. (1987) A conflict between two sexes, females and hermaphrodites. In S.C. Stearns (ed.) The Evolution of Sex and its Consequences. Birkhaeuser Verlag, Basel, pp. 245-261. Hamilton, W.D. (1967) Extraordinary sex ratios. Science 156, 477-488. Hamilton, W.D. and May, R.M. (1977) Dispersal in stable habitats. Nature 269, 578-581. Holsinger, K.E. (1986) Dispersal and plant mating systems: the evolution of self-fertilization in subdivided populations. Evolution 40, 405-413. Holsinger, K.E. (1988) Inbreeding depression doesn't matter: the genetic basis of mating-system evolution. Evolution 42, 1235-1244. Holsinger, K.E. (1992) Ecological models of plant mating systems and the evolutionary stability of mixed mating systems. In R. Wyatt (ed.) Ecology and Evolution of Plant Reproduction. Chapman and Hall, New York, pp. 169-191. Ishida, K. and Hiura, T. (1998) Pollen fertility and flowering phenology in an androdioecious tree, Fraxinus lanuginosa (Oleaceae), in Hokkaido, Japan. Int. J. Pl. Sci. 159, 941-947. Johnston, M.O. and Schoen, D.J. (1996) Correlated evolution of self-fertilization and inbreeding depression: an experimental study of nine populations of Amsinckia (Boraginaceae). Evolution 50, 1478-1491. Karron, J.D., Thumser, N.N., Tucker, R. and Hessenauer, A.J. (1995) The influence of population density on outcrossing rates in Mimulus ringens. Heredity 75, 175-180. Kennington, W.J. and James, S.H. (1997) The effect of small population size on the mating system of a rare clonal mallee, Eucalyptus argutifolia (Myrtaceae). Heredity 78, 252-260. Kohn, J.R. and Barrett, S.C.H. (1994) Pollen discounting and the spread of a selfing variant in tristylous Eichhornia paniculata: evidence from experimental populations. Evolution 48, 1576-1594. Lande, R. and Schemske, D.W. (1985) The evolution of self-fertilization and inbreeding depression. Evolution 39, 24-40. Lavorel, S., O'Neil, R.V. and Gardner, R.H. (1994) Spatio-temporal dispersal strategies and annual plant species in a structured landscape. Oikos 71, 75-88. Lemen, C. (1980) Allocation of reproductive effort to the male and female strategies in wind-pollinated plants. Oecologia 45, 156-159. Lepart, J. and Dommee, B. (1992) Is Phillyrea angustifolia L. (Oleaceae) an androdioecious species? Bot. J. Linn. Soc. 108, 375-387. Levin, D.A. (1996) The evolutionary significance of pseudo-self-fertility. Am. Nat. 148, 321-332. Lipow, S.R., Broyles, S.B. and Wyatt, R. (1999) Population differences in self-fertility in the “self-incompatible” milkweed Asclepias exaltata (Asclepiadaceae). Am. J. Bot. 86, 1114-1120. Liston, A., Rieseberg, L.H. and Elias, T.S. (1990) Functional androdioecy in the flowering plant Datisca glomerata. Nature 343, 641-642. Lloyd, D. (1975) The maintenance of gynodioecy and androdioecy in angiosperms. Genetica 45, 325-339. Lloyd, D.G. (1965) Evolution of self-compatibility and racial differentiation in Leavenworthia (Cruciferae). Contributions to the Gray Herbarium 195, 3-134. Lloyd, D.G. (1974) The genetic contributions of individual males and females in dioecious and gynodioecious angiosperms. Heredity 32, 45-51. Lloyd, D.G. (1982) Selection of combined versus separate sexes in seed plants. Am. Nat. 120, 571-585. Lloyd, D.G. (1987) Allocations to pollen, seeds and pollination mechanisms in self-fertilizing plants. Funct. Ecol. 1, 83-89. Lloyd, D.G. and Bawa, K.S. (1984) Modification of the gender of seed plants in varying conditions. Evol. Biol. 17, 255-336. Lloyd, D.G. and Schoen, D.J. (1992) Self-and cross-fertilization in plants. I. Functional dimensions. Int. J. Pl. Sci. 153, 358-369. Lu, A. and Zhang, A. (1986) Schizopepon. In A. Lu and S. Chen (eds) Flora Republicae Popularis Sinicae. Science Press, Beijing, pp. 181-188. Manicacci, D., Atlan, A. and Couvet, D. (1997) Spatial structure of nuclear factors involved in sex determination in the gynodioecious Thymus vulgaris L. J. Evol. Biol. 10, 889-907. Mayer, S.S. and Charlesworth, D. (1991) Cryptic dioecy in flowering plants. Trends Ecol. Evol. 6, 320-325. Olivieri, I., Couvet, D. and Gouyon, P.H. (1990) The genetics of transient populations: research at the metapopulation level. Trends Ecol. Evol. 5, 207-210. Olivieri, I., Gouyon, P.H. and Gilpin, M.E. (1997) Evolution of migration rate and other traits: the metapopulation effect. In I. Hanski (ed.) Metapopulation Biology: Ecology, Genetics, and Evolution. Academic Press, San Diego, pp. 293-323. Otto, S.P., Sassaman, C. and Feldman, M.W. (1993) Evolution of sex determination in the conchostracan shrimp Eulimnadia texana. Am. Nat. 141, 327-337. Pannell, J. (1997a) The maintenance of gynodioecy and androdioecy in a metapopulation. Evolution 51, 10-20. Pannell, J. (1997b) Variation in sex ratios and sex allocation in androdioecious Mercurialis annua. J. Ecol. 85, 57-69. Pannell, J (1997c) Widespread functional androdioecy in Mercurialis annua L. (Euphorbiaceae). Biol. J. Linn. Soc. 61, 95-116. Pannell, J.R. and Barrett, S.C.H. (1998) Baker's Law revisited: reproductive assurance in a metapopulation. Evolution 52, 657-668. Pannell, J.R. and Barrett, S.C.H. (2001) Effects of population size and metapopulation dynamics on a mating-system polymorphism. Theor. Pop. Biol. (in press). Pannell, J.R. and Ojeda, F. (2000) Patterns of flowering and sex-ratio variation in the Mediterranean shrub Phillyrea angustifolia (Oleaceae): implications for the maintenance of males with hermaphrodites. Ecol. Let. 3, 495-502. Ramsey, M. (1993) Floral morphology, biology and sex allocation in disjunct populations of Christmas bells (Blandfordia grandiflora, Liliaceae) with different breeding systems. Aus. J. Bot. 41, 749-762. Rieseberg, L.H., Hanson, M.A. and Philbrick, C.T. (1992) Androdioecy is derived from dioecy in Datiscaceae: evidence from restriction site mapping of PCR-amplified chloroplast DNA fragments. Syst. Bot. 17, 324-336. Ritland, C.R. and Ritland, K. (1989) Variation of sex allocation among eight taxa of the Mimulus guttatus species complex (Scrophulariaceae). Am. J. Bot. 76, 1731-1739. Ross, M.D. (1982) Five evolutionary pathways to subdioecy. Am. Nat. 119, 297-318. Routley, M.B., Mavraganis, K. and Eckert, C.G. (1999) Effect of population size on the mating system in a self-compatible, autogamous plant, Aquilegia canadensis (Ranunculaceae). Heredity 82, 518-528. Sassaman, C. (1991) Sex ratio variation in female-biased populations of Notostracans. Hydrobiologia 212, 169-179. Sassaman, C. and Weeks, S.C. (1993) The genetic mechanism of sex determination in the conchostracan shrimp Eulimnadia texana. Am. Nat. 141, 314-328. Schoen, D.J. (1982) The breeding system of Gilia achilleifoliaI: variation in floral characteristics and outcrossing rate. Evolution 36, 352-360. Stebbins, G.L. (1957) Self-fertilization and population variability in the higher plants. Am. Nat. 91, 337-354. Stephenson, A.G., Good, S.V. and Vogler, D.W. (2000) Interrelationships among inbreeding depression, plasticity in the self-incompatibility system, and the breeding system of Campanula rapunculoides L-(Campanulaceae). Ann. Bot. 85, 211-219. Swensen, S.M., Luthi, J.N. and Rieseberg, L.H. (1998) Datiscaceae revisited: monophyly and the sequence of breeding system evolution. Syst. Bot. 23, 157-169. Taylor, D.R., Trimble, S. and McCauley, D.E. (1999) Ecological genetics of gynodioecy in Silene vulgaris: relative fitness of females and hermaphrodites during the colonization process. Evolution 53, 745-751. Tonsor, S.J. (1985) Leptokurtic pollen-flow, non leptokurtic gene-flow in a wind-pollinated herb, Plantago lanceolata L. Oecologia 67, 442-446. Traveset, A. (1994) Reproductive biology of Phillyrea angustifolia L. (Oleaceae) and effect of galling-insects on its reproductive output. Bot. J. Linn. Soc. 114, 153-166. Tutin, T.G., Heywood, V.H., Burges, N.A., Moore, D.M., Valentine, D.H., Walters, S.M. and Webb, D.A. (1968) Flora Europaea. Cambridge University Press, Cambridge. Vantreuren, R., Bijlsma, R., Ouborg, N.J. and Vandelden, W. (1993) The effects of population size and plant density on outcrossing rates in locally endangered Salvia pratensis. Evolution 47, 1094-1104. Vassiliadis, C., Lepart, J., Saumitou-Laprade, P. and Vernet, P. (2000) Self-incompatibility and male fertilisation success in Phillyrea angustifolia (Oleaceae). Int. J. Pl. Sci. 161, 393-402. Webb, C.J. (1979) Breeding systems and the evolution of dioecy in New Zealand apioid Umbelliferae. Evolution 33, 662-667.